1. travel
2. tourist destinations
3. australia and new zealand

© Copyright Australian Museum, 2002
Records of the Australian Museum (2002) Vol. 54: 313–324. ISSN 0067-1975
Bandella, A New Hilarine Fly Genus from Australia
(Diptera: Empididae)
D ANIEL J. B ICKEL
Australian Museum, 6 College Street, Sydney NSW 2010, Australia
[email protected]
ABSTRACT. Bandella n.gen. (Diptera: Empididae: Empidinae: Hilarini) is described from mainland
Australia and Tasmania, and comprises nine new species: B. albitarsis, B. cerra, B. allynensis, B. costalis,
B. duvalli, B. maxi, B. montana, B. noorinbee, and B. tasmanica. Bandella is distinguished from other
genera in the Hilarini by the following combination of characters: mesonotum glabrous, without
pruinosity, but with diagnostic colour patterns; mesonotal vestiture highly reduced, comprising only
short setulae; male tarsus I unmodified; and male abdominal terga 7 and 8 reduced or modified to form
flaps so that hypopygium can be flexed forward. The genus is distinctive in having an almost fully
formed vein CuP running lengthwise across the centre of cell cup. Although vein CuP is part of the wing
groundplan in Diptera, it has variable expression in the lower Brachycera. In other Empididae, this vein
is either residual and positioned closely posteriad of vein CuA2, or totally absent. Also, Bandella has a
divided and sclerotised male cercus, a character also shared by the endemic Tasmanian hilarine genus
Cunomyia Bickel. Bandella has a predominately temperate Bassian distribution: Tasmania, southeastern
and southwestern Australia, with an outlying species in submontane rainforest of tropical Queensland.
BICKEL, DANIEL J., 2002. Bandella, a new hilarine fly genus from Australia (Diptera: Empididae). Records of the
Australian Museum 54(3): 313–324.
This is the third contribution to a revision of the Australian
Hilarini (Diptera: Empididae: Empidinae) (see Bickel, 1996,
1998). This tribe is taxonomically rich in Australia, with
hundreds of species awaiting description, especially in the
complex of genera similar to the cosmopolitan genus Hilara.
This short paper describes Bandella, a distinctive new
genus comprising nine new species: eight are from
temperate Australia and one is from submontane rainforest
in the Queensland wet tropics. It is distinguished by a
strongly developed vein CuP, a thorax almost totally devoid
of major setae but with shining patterned cuticle, an
unswollen (i.e., unmodified) male basitarsis I, and by a
tendency to curl the distal tarsomeres on all legs (dried
specimens). Bandella is morphologically isolated with
respect to other genera in the Hilarini, although it possibly
is closest to the Tasmanian hilarine genus Cunomyia Bickel,
as both genera share a divided male cercus.
Materials and methods
This study is based on material housed in major Australian
and overseas collections (see Acknowledgements for list
of repositories and their abbreviations). Morphological
terminology follows McAlpine (1981) except that of the
male terminalia, which follows Cumming et al. (1995).
Measurements are in millimetres and based on representative dry specimens. The position of features on elongate
structures such as leg segments is given as a fraction of the
total length, starting from the base. The relative lengths of
the podomeres are representative ratios and not measurements, and are presented in the following formula and
punctuation: trochanter + femur; tibia; tarsomere 1/ 2/ 3/ 4/ 5.
The following abbreviations and terms are used
(museums and other collections are listed in Acknowledgments): FSSC, female secondary sexual character(s), the
www.amonline.net.au/pdf/publications/1369_complete.pdf
314
Records of the Australian Museum (2002) Vol. 54
non-genitalic characters found only on female body; MSSC,
male secondary sexual character(s), the non-genitalic
characters found only on male body; I, II, III: pro-, meso-,
metathoracic legs; C, coxa; T, tibia; F, femur; ac, acrostichal
setae; ad, anterodorsal; av, anteroventral; dc, dorsocentral
setae; dv, dorsoventral; pd, posterodorsal; pv, posteroventral; t, tarsus; t1–5, tarsomeres 1 to 5. Geographical
abbreviations: FR, Flora Reserve; NP, National Park; NR,
Nature Reserve; SF, State Forest.
Genus Bandella n.gen.
Type species. Bandella allynensis n.sp.
Diagnosis. Flies in the tribe Hilarini; mesonotum glabrous,
without pruinosity, but with species diagnostic colour
patterns present; mesonotum with ac and dc reduced to
setulae, and posterior slope covered with field of scattered
setulae; tibia I without distinct anteroapical comb; male
basitarsus not swollen; costa circumambient, although
reduced in thickness along posterior margin; Sc incomplete;
R4+5 branched, with R4 diverging at 45° angle and slightly
sinuate, and R5 ending behind wing apex; CuA2, strongly
recurrent; vein CuP present as distinct vein across cell cup;
abdomen terga without strong marginal setae; tergum 7 often
with U-shaped median excavation; tergum 8 usually with
lateral flaplike projections; hypopygium can flex anteriad
at rest; male cercus divided into short basal plate and
digitiform clasping cercus.
Description. Rather large, body length 7.0–11.5.
Head spheroidal, with convex post-cranium; post-cranium
sometimes with distinctive pruinosity, vertex and frons
usually shining with little pruinosity; major setae absent;
postcranium with scattered pale ventral hairs, postorbitals
short; postvertical setae very short, black; ocellar triangle
not distinctly raised; ocellar setae reduced to short hairs;
frons slightly narrower than ocellar triangle, and with very
short hairs along lateral margins; eyes notched laterad of
antennae; face as wide as frons and slightly expanded
apically, often covered with pruinosity; palp yellow with
white setae, curved and slightly clavate; labrum tapering
and heavily sclerotised, about one and a half head height in
length, and projecting ventrally; labellum separated from
and subequal with labrum; antenna (Fig. 3c) scape about
twice as long as pedicel; first flagellomere tapering, longer
than scape and pedicel combined, and with short 2–articled
apical style.
Thorax. Mesonotum glabrous, without pruinosity, but
pleura with pruinosity; mesonotum with species diagnostic
colour patterns present, usually similar in both sexes, but
sometimes sexually dimorphic; mesonotum with only short
setulae and apart from following major setae: 2 posterior
notopleural (npl) setae, 1 short postalar (pa) seta, and 1 short
posterior intra-alar seta: ac band 3–4 setulae wide, with bare
cuticle each side of this band; remainder of mesonotum,
including posterior slope covered with field of scattered
short setulae; prothoracic collar (pronotum) with some short
setae; proepisternum with field of 8–10 pale hairs; scutellum
rather weakly developed and with pair of median marginal
setae and some adjacent short lateral setae; laterotergite bare;
postnotum broad and bulging.
Legs. All tarsi with strong black claws and large yellowish
pulvilli; tarsi often curled in repose in dried specimens; FI
and TI mostly with short vestiture; TI without distinct
anteroapical comb, although some species have irregular
setal row of variable length; male It1 not swollen; TI and
each tarsomere It1–3 with 2–3 pairs of subapical av-pv setae,
and It1–3 with dense ventral vestiture; TII with pair strong
av-pv setae; each tarsomere IIt1–3 with apical setae; IIt2–5
each slightly flattened with pale ventral pile; FIII weakly
expanded in distal third; TIII without strong setae.
Wing (Fig. 3b); membrane hyaline; costa circumambient,
although reduced in thickness along posterior margin; Sc
distinctly incomplete; R1 slightly swollen near join with
costa, and brownish stigma present under distal R1; costa
haired, but other veins bare; R 4+5 branched, with R 4
diverging at 45° and slightly sinuate, and R5 ending behind
wing apex; M1, M2, and CuA1 all joining margin; CuA2,
which closes cell cup, strongly recurrent; vein CuP present
as almost fully formed vein across cell cup; A1 present
distally only as fold, and arising midway along cell cup; A2
present as trace; anal angle weak to absent; lower calypter
yellowish with yellow setae; halter yellow.
Abdomen. Cuticle glabrous, with little pruinosity; tergum
1 with some pale setae laterally, otherwise abdomen with
only short yellowish vestiture, without strong marginal
setae; tergum 1 relatively short with membranous median
window; abdominal plaques present on terga 2–5; tergum
7 often with U-shaped median excavation; tergum 8 usually
with lateral flaplike projections (e.g., 4b,c); hypopygium
can flex anteriad at rest; male cercus distinct and divided
into sclerotised short basal cercal plate and digitiform
clasping cercus; hypandrium keel-like; aedeagus elongate
and conforming to curvature of hypandrium, epandrium
broad, with distal curved projection; distinct surstylus not
evident.
Female oviscapt relatively unmodified, with subequal terga
and sterna on segments 9 and 10, and with pair elongate
apical cerci (Fig. 3f).
Etymology. Bandella means “little Banda”, a diminutive
derived from Mt Banda Banda, a New South Wales locality
where the genus occurs. The gender is feminine.
Remarks
Bandella comprises nine species from mainland Australia
and Tasmania. Species occur mostly in wet sclerophyll
eucalypt forests and rainforests, and mostly at higher
altitudes. Although the genus has an essentially temperate
Australian or Bassian distribution of southeastern and
southwestern Australia and Tasmania, one species occurs
above 1000 m in submontane rainforest in tropical
Queensland.
Bandella is best placed in the Hilarini, as discussed in
the next section. It is confined to Australia, and is not close
to any described genus from temperate Gondwanan regions:
New Zealand (Collin, 1928), Patagonia (Collin, 1933) and
southern Africa (Smith, 1969). However, based on a shared
character, the divided sclerotised male cercus, Bandella
possibly has a distant relationship with the endemic
Tasmanian genus Cunomyia Bickel.
Bickel: a new empidid fly genus
Bandella has a glabrous thorax with strong colour
patterns, and almost all major dorsal thoracic setae are
reduced to setulae. By contrast, most other hilarines have a
pruinose thoracic surface, and at least some strong thoracic
setae, especially the anterior dc. Variation is evident in the
thoracic pattern of some Bandella species, which may reflect
age, nature of preservation or intraspecific variation. For
example, the strong contrast between yellow and dark brown
on some specimens may be obscured by a general
infuscation of the entire specimen, possibly a direct function
of age (e.g., Figs. 2h,i). Although the thoracic patterns are
usually the same in both sexes, they are sometimes strikingly
dimorphic, as in B. noorinbee (Fig. 2d,e). However, the basic
contrasting thoracic pattern as suggested by the figures
remains diagnostic for each species and is useful in
identifying females.
The male postabdomen is variously modified to receive
the deflexed hypopygium. In all species except B. maxi
tergum 7 is medially excavated, and in some species tergum
8 has lateral flap-like projections (Fig. 4b,c). In dried
specimens of Bandella, particularly B. costalis, B. duvalli,
B. montana, and B. tasmanica, the tarsi are distinctly curled
in repose, as in Fig. 1.
315
There is good evidence of adult nectar feeding in the
genus. Bandella duvalli was collected off the nectar-rich
blossoms of Leptospermum (Myrtaceae) bushes in
Tasmania, and Leptospermum pollen was found on the
Figure 1. Bandella duvalli, 웧, habitus. Mt Pelion, Tasmania.
316
Records of the Australian Museum (2002) Vol. 54
Figure 2. Dorsal thoracic colour pattern, Bandella spp. a, B. allynensis 웧; b, B. cerra 웧; c, B. montana 웧; d, B.
noorinbee 웧; e, B. noorinbee 웨; f, B. albitarsis 웧; g, B. maxi 웧; h, B. duvalli 웧 (holotype); i, B. duvalli 웧
(variant); j, B. tasmanica 웧; k, B. costalis 웧.
proboscis of B. albitarsis in New South Wales. Nectar
feeding is well known among the Empidinae (e.g., Chvála,
1976). Specimens of B. allynensis have also been taken on
sticky traps placed on smooth barked Eucalyptus trunks,
which indicates adults occasionally rest on tree trunks.
Mating behaviour is unknown. Males do not have
swollen fore basitarsi producing silk for wrapping nuptial
gifts characteristic of the complex of genera near Hilara.
Of particular interest are the bright silvery pruinose patches
present on the head, thorax and legs of both sexes of
Bandella albitarsis. These silvery patches may facilitate
sexual recognition in flight, especially since incident light
would make them appear to “flash” while turning.
Morphological notes and
systematic position of Bandella
I presented (Bickel, 1996) a set of characters useful in
separating the two major tribes of the Empidinae, the
Empidini and the Hilarini. Since many characters are
variable in expression and must be qualified, higher level
taxa have to be defined as a mosaic of characters, a
“polythetic classification,” no character necessarily being
diagnostic for all members (Gauld & Mound, 1982). As so
defined, Bandella has the following set of characters that place
it in the Hilarini: laterotergite bare; hypandrium forming a
curved convex hood over aedeagus; costa circumambient; vein
R1 distinctly swollen before it joins the costa. Bandella does
not have the following hilarine characters: tibia I with
anteroapical comb of 8–10 short even setae; male cercus small
Bickel: a new empidid fly genus
and desclerotised, and fused laterally with the surstylus and
epandrium; male basitarsus I enlarged or swollen.
With the large number of undescribed taxa in the Hilarini,
both in Australia and throughout the world, it is premature
to attempt any phylogenetic analysis of the tribe. However,
two important morphological characters should be
discussed—vein CuP and the male cercus.
Vein CuP. Bandella is distinctive in having an almost
fully formed vein CuP running lengthwise across the centre
of cell cup (Fig. 3b), and not posteriorly close to vein CuA2
or vestigial as found in some other empidids (e.g.,
Brachystoma and some Empis, figures in Chvála, 1983).
However vein CuP is absent in Hilara, most Empis spp.
and the Trichopezinae. By contrast in Bandella, CuP is
strongly developed in all species, although it looks almost
like a vena spuria, and not carrying haemolymph. Vein CuP
is plesiomorphic to the Diptera as a whole and part ground
plan for the Diptera wing (McAlpine, 1981), but its
expression is variable in the lower Brachycera. For example,
it is variously developed in the Asiloidea, but usually weak
and just behind vein CuA2.
Cercus. Most male Hilarini have a cercus which is rather
small, desclerotised, and fused laterally with the surstylus
and epandrium (Cumming & Sinclair, unpubl. data).
Bandella, by contrast, has an enlarged sclerotised male
cercus, which is divided into a distinctive digitiform
“clasping cercus” and smaller “cercal plate” (see Fig. 4b).
This cercal form is similar to that of the monotypic
Tasmanian genus Cunomyia Bickel (see Sinclair, 1995 and
Bickel, 1998 for further discussion), and this shared
character indicates a possible close phylogenetic relationship between these two Australian genera.
Key to species of Bandella (males)
1
Abdomen almost entirely shining black with violet reflections,
with at most some lateral yellow colour on tergum 1 ................................................................... 2
—— Abdomen with some red, brown or yellow terga .......................................................................... 3
2
Large, length >10.0; thoracic pattern, mesonotum mostly orange,
but black over posterior slope, scutellum mostly orange, Fig. 2c;
legs mostly yellow; hypopygium (Fig. 3d) mostly yellow (NSW,
Vic.) ...................................................................................................................... B. montana n.sp.
—— Small, length <7.5; male mesonotum and scutellum black, Fig.
2d (female thorax completely orange, Fig. 2e); scutellum black;
coxae yellow; remainder of legs dark brown to black; hypopygium
(Fig. 3e) black (Vic., NSW) ............................................................................... B. noorinbee n.sp.
3
Post-cranium with silvery pruinose band around outer rim; thoracic
pattern, Fig. 2f; pleuron with bright silvery pruinose areas; It2–5
white; IIIt2–5 pale yellow; abdomen, including hypopygium mostly
yellow (NSW) ..................................................................................................... B. albitarsis n.sp.
—— Head and thorax without silvery pruinosity; tarsus I yellow to
black, but not white ........................................................................................................................ 4
4
TII and tarsus II with long anterior and posterior setae, Fig. 5e;
mesonotum and scutellum almost entirely black, Fig. 2k; FIII
slightly clavate in distal half; abdominal terga yellow with black
posterior border (WA) ............................................................................................ B. costalis n.sp.
—— Legs without long setae, only with normal short vestiture;
mesonotum with some red, yellow or brown colouration; FIII and
abdomen variously developed ....................................................................................................... 5
5
TIII banded, with yellow apex and base, and dark brown median
band (as in Fig. 1) (Tas.) ................................................................................................................ 6
—— TIII either entirely yellow or yellow with dark brown apex (Aust.
mainland) ........................................................................................................................................ 7
6
317
Coxae dark brown/black; anterior mesonotum mostly red-brown;
posterior mesonotum black with interdigitation of red anteriorly
(Fig. 2h,i); FIII yellowish in basal half but distally dark brown;
tarsus III dark brown; hypopygium (Fig. 4b,c) mostly black ................................ B. duvalli n.sp.
—— Coxae mostly yellow; anterior mesonotum mostly orange; with
black ac band and black spot laterad of dc row (Fig. 2j); FIII mostly
red-yellow; tarsus III entirely yellow; hypopygium (Fig. 4a) mostly
yellow except for dark brown apex ................................................................... B. tasmanica n.sp.
318
Records of the Australian Museum (2002) Vol. 54
7
Antenna entirely dark brown; anterior mesonotum red-brown,
interdigitating with black posterior mesonotum (Fig. 2g);
scutellum black with reddish margin; postnotum black; abdominal
terga 1 and 2 reddish, hypopygium (Fig. 4d) with hypandrium
greatly expanded and keel-like (n. Qld) ..................................................................... B. maxi n.sp.
—— Antennal scape and pedicel yellowish, first flagellomere black;
pleura and postnotum dark brown with yellowish areas; anterior
mesonotum mostly yellow; postpronotum and postalar callus pale
yellow; scutellum yellow but infuscated basally; hypopygium not
expanded, similar to Fig. 3a ........................................................................................................... 8
8
Coxae II and III dark brown; thorax with posterior slope of
mesonotum dark brown and with thin brown strip extending
anteriorly over dc band (Fig. 2a); TIII mostly yellow; It1 yellow;
hypopygium (Fig. 3a) entirely dark brown (NSW) ........................................... B. allynensis n.sp.
—— Coxae II and III mostly yellow; mesonotum with three brown
bands extending to scutellum and with lateral dark brown areas
(Fig. 2f); TIII yellow but with distal fifth brown; It1 distinctly
brown, in contrast to yellow TI and tarsomeres It2–4; hypopygium
with dark brown hypandrium and surstyli, but distinctly yellow
epandrium (NSW) ...................................................................................................... B. cerra n.sp.
Bandella allynensis n.sp.
Figs. 2a, 3a–c
Type material. HOLOTYPE 웧 and PARATYPES, 2웧, 10웨 all
New South Wales, Upper Allyn River, 1500 ft (410 m),
9.xi.1960, I.F.B. Common & M.S. Upton (ANIC).
Additional material. New South Wales: 3웧, 2웨, New
England NP, Point Lookout, 1400 m, Nothofagus forest,
22.i.1967 (UQIC); Wilson River Reserve, 31°12'S 152°28'E,
240 m, wet sclerophyll/ subtropical rainforest: 웨,
26.xi.1966, & 웧, sticky trap on Eucalyptus grandis, 16–
19.xi.1998; 웧, 웨 Carrai SF, Daisy Plains, 30°54'19"S
152°17'36"E, 1055 m, 11–16.i.1998, wet sclerophyll forest,
sticky trap on eucalyptus trunk (AMS).
Description (male): length: 9.4–9.8; wing: 8.2×1.4. Head:
post-cranium black, covered laterally and ventrally with
grey pruinosity; dorsal post-cranium, vertex and frons black
without pruinosity; postcranium with usual pale ventral
hairs; dorsal postcranium bare; postorbital setae white
ventrally, black dorsally; labrum dark brown but yellow at
middle; labellum dark brown; antennal scape and pedicel
yellowish to brown; first flagellomere black.
Thorax: pleura and postnotum dark brown with yellowish
areas, and covered with grey pruinosity; anterior mesonotum
mostly yellow with pale yellow postpronotum and postalar
callus; ac band dark brown, and dark brown stripe present
from postpronotum to postalar callus; posterior slope of
mesonotum dark brown, with thin brown strip extending
anteriorly over dc band, but fading out near mesonotal suture
(Fig. 2a); scutellum yellow but infuscated at base; posterior
slope of mesonotum with some short setulae.
Legs: CI yellow and trochanter I yellow with dark brown
apical rim; coxae and trochanters II and III distinctly dark
brown; remainder of leg I yellow except distal tarsomeres
infuscated; FII and FIII basally yellow; but infuscated
distally; TII and TIII mostly yellow, but TIII infuscated
apically; tarsi II and III brown; leg vestiture brownish; coxae
with only short pale vestiture, no strong setae; all tarsi with
strong black claws and large yellowish pulvilli; I: 11.0; 11.5;
6.0/ 3.5/ 1.5/ 1.2/ 1.3; FI and TI mostly with short vestiture;
It2 and It3 each with two closely appressed subapical pv
setae, which are distinctly longer than single corresponding
single av seta.(MSSC); It2–5 ventrally flattened with pale
vestiture; II: 11.0; 11.5; 6.0/ 3.5/ 1.5/ 1.2/ 1.3; TII with pair
strong av-pv setae; each tarsomere IIt1–3 with apical setae;
IIt2–5 each slightly flattened with pale ventral pile; III: 15.0;
17.0; 7.0/ 3.0/ 1.3/ 1.8/ 1.6; FIII weakly expanded in distal
third; TIII without strong setae; IIIt with short black
vestiture.
Abdomen: tergum 1 dark brown; tergum 2 with basal
third yellow and distally dark brown; terga 3–6 mostly
yellow but each tergum with dark brown posterior marginal
band (in darker specimens, marginal band larger and with
dorsoanterior extensions on most terga); cuticle glabrous,
without pruinosity; tergum 7 with U-shaped median
excavation; tergum 8 unmodified; hypopygium (Fig. 3a);
male cercus divided into short basal cercal plate and
digitiform clasping cercus.
Female similar to male except as noted: thorax with similar
colour; leg vestiture somewhat weaker than on male; It2
and It3 each with single subequal subapical av-pv setal pair.
Remarks. Bandella allynensis occurs in subtropical and
temperate rainforests and wet sclerophyll eucalypt forests
along the ranges of northeastern New South Wales.
Specimens taken in sticky traps (sheets of clear plastic
coated with sticky substance and attached to tree trunks)
suggest the species occasionally rests on tree trunks. There
is some variation in the extent of the dark brown area on
the mesonotum (Fig. 2a).
Bandella cerra n.sp.
Fig. 2b
Type material. HOLOTYPE, 웧 (K 173134), PARATYPES, 웧
(K 173135), 웨 (K 173136), New South Wales: Barrington
Tops NP, Gloucester Tops, 32°04'S 151°34'E, 1280 m,
Nothofagus moorei forest along creek, 4–30.xii.1988,
malaise trap, D.J. Bickel (all AMS).
Bickel: a new empidid fly genus
319
Figure 3. Bandella allynensis: a, male postabdomen; b, male wing, dorsal; c, male antenna, left lateral. Bandella
montana: d, male postabdomen, left lateral. Bandella noorinbee: e, male postabdomen, left lateral; f, female oviscapt,
dorsal.
Additional material. New South Wales: 웧, Barrington
Tops, via Salisbury [no elevation], 28–30.xii.1965 (UQIC);
웧, Mt Banda Banda FR, 31°10'S 152°26'E, 1200 m, cool
temperate rainforest, 14.i.1988 (AMS).
Description (male): length: 8.5–9.5; wing: 6.9×1.3; similar
to B. allynensis except as noted. Head: scape black. Thorax:
pleura and postnotum dark brown with yellowish areas, and
covered with grey pruinosity; anterior mesonotum mostly
yellow with pale yellow postpronotum and postalar callus;
ac band dark brown and extending to scutellum, with
yellow bands on either side, also extending to scutellum,
and with brown areas laterad which extend from
mesonotal suture back to scutellum (Fig. 2b); scutellum
yellow but infuscated at base.
Legs: CI yellow; trochanter I yellow with dark brown
apical rim; coxae and trochanters II and III yellowish with
brown areas, and covered with grey pruinosity; FI and TI
320
Records of the Australian Museum (2002) Vol. 54
yellow; It1 distinctly brown, in contrast to yellow TI and
distal tarsomeres; It2–5 yellow; It2 and It3 each with two
closely appressed subapical pv setae, which are distinctly
longer than single corresponding single av seta.(MSSC);
It2–5 ventrally flattened with pale vestiture; FII and TII
yellow; It1–2 brown, and It3–5 yellow; FIII yellow basally,
grading to brown in distal third; TIII yellow with but distal
quarter brown; IIIt dark brown.
Abdomen: tergum 1 dark brown; terga 2–6 mostly yellow
but each tergum with dark brown apical ring and with dark
brown dorsal band so that abdomen appears to have dorsal
stripe; hypopygium (not figured, but similar to Fig. 3a) with
dark brown hypandrium and surstyli, but distinctly yellow
epandrium.
with dense ventral vestiture, and with some subapical avpv setae; III: 18.0; 17.0; 6.3/ 2.1/ 1.8/ 1.5/ 1.8; FIII very
slightly clavate in distal third; tarsomeres IIIt1–2 ventrally
swollen with dense vestiture.
Abdomen: segments 1–6 shining black with some violet
reflections, although tergum 2 yellow in basolaterally;
segments 7 and 8 yellowish; tergum 6 dorsally with medianposterior U-shaped excavation, and tergum 7 with lateral
flaplike projections; hypopygium yellowish (Fig. 3d).
Female similar to male except as noted: thorax with similar
colour; leg vestiture somewhat weaker than on male; It2
and It3 each with single subequal subapical av-pv setal pair;
abdomen with similar pattern.
Remarks. Bandella montana occurs along the Great
Dividing Range from southern Victoria to the New England
escarpment in northern New South Wales. The extent of
the black area on the mesonotum appears to vary in size
geographically. Specimens from the southern part of the
range (Blue Mountains, N.S.W. and Thompson River, Vic.)
have the black area reduced to a narrow median strip,
whereas specimens from the New England escarpment have
a more extensive black area, as in Fig. 2c.
Remarks. Bandella cerra is known from Nothofagus
moorei dominated cool temperate rainforest in the
Barrington Tops and Mt Banda Banda, in northeastern New
South Wales. These sites are in the Central Eastern
Rainforest Reserves of Australia (CERRA) World Heritage
Area, hence the specific epithet.
Female similar to male except as noted: post-cranium also
with grey pruinosity; thorax with similar colour and black
patch size variation; leg vestiture somewhat weaker than
on male; abdominal tergum 1 with similar colour pattern.
Bandella montana n.sp.
Bandella noorinbee n.sp.
Figs. 2c, 3d
Figs. 2d, 2e, 3e, 3f
Type material. HOLOTYPE, 웧 (K 173137), PARATYPES, 웧
(K 173138), 웨 (K 173139), New South Wales: Barrington
Tops NP, 5.xii.1976, G. Daniels (all AMS).
Type material. H OLOTYPE 웧, PARATYPES 2웧, 11웨,
Victoria: Noorinbee, 12.xi.1969, A. Neboiss (all MVM).
Additional material. NEW SOUTH WALES : NSW: Mt
Wilson, Blue Mountains, 33°30'S 150°23'E, 950 m, 웨,
5.xii.1956, 웨, 1.xi.1975, 웧, 10.xi.1990 (AMS); 웧, New
England NP, 5.xi.1981 (AMS); 웧,Mt Tomah, Blue Mtns,
33°33'S 150°25'E, x.1930 (MVM). VICTORIA: 웨, Thomson
River, 6 km E of Mt Gregory, 24.xi.1976 (MVM).
Description (male): length: 11.0; wing: 8.8×2.7. Head: postcranium laterally and ventrally with dusting of grey
pruinosity; dorsal postcranium, vertex and frons shining
black with no pruinosity; ventral postcranium with scattered
pale setae; postorbitals present as short white setae ventrally,
black dorsally; face covered with silvery pruinosity; labrum
and labellum entirely yellow; antenna black but pedicel
brownish in some specimens.
Thorax: pleura and mesonotum mostly red-yellow, except
posterior mesonotum with black area extending almost to
scutellum, but varying in extent from large area (Fig. 2c) to
a small black dot, almost absent; scutellum yellow but
brownish basally; pleura with some pruinosity, but
mesonotum glabrous; scutellum with pair brownish median
marginal setae, with some adjacent short setae.
Legs: coxae and remainder of legs yellow; coxae with
only short pale vestiture, no strong setae; femora and tibiae
with mostly short black vestiture; all tarsi with strong black
claws and large yellowish pulvilli; I: 11.5; 11.5; 5.0/ 2.0/
1.6/ 1.4/ 1.5; TI and each tarsomere It1–3 with 2–3 pairs of
subapical av-pv setae, and It1–3 with dense ventral vestiture;
II: 12.5; 12.0; 5.0/ 2.0/ 1.2/ 1.1/ 1.5; each tarsomere IIt1–4
Additional material. New South Wales: 3웧, 6.4 km N of
Batemans Bay, 14.x.1952, 22.x.1952, 14.x.1959; 웧,
Macquarie Falls, 14.xi.1960, 2웨, Warrell Creek, 11.x.1962
(ANIC). 웧, Mt Royal SF, nr Pieries Ridge, 600–700 m,
15.xi.1986; 웨, 30 km N of Taree, 23.xi.1985 (AMS); 웧,
Royal NP, Couranga Trail, 7.xi.1993 (CNC).
Description (male): length: 7.2–7.3; wing: 5.0×1.5. Head:
post-cranium, vertex and frons shining black with no
pruinosity; ventral postcranium with scattered pale setae;
postorbitals reduced to short hairs, white ventrally, black
dorsally; face covered with silvery pruinosity; labrum
yellowish with dark brown base and apex, labellum black;
antenna black brown; scape and pedicel setose.
Thorax: pleura and humeral areas of mesonotum redyellow; mesonotum mostly black posteriorly, narrowing step
like anteriorly, to become black stripe over ac band and
pronotum (Fig. 2d); scutellum black; postnotum black but
covered with grey pruinosity; setulae pale; scutellum with
pair short median marginal setae, and with some adjacent
short setae.
Legs: coxae yellow; trochanters and remainder of legs
dark brown to black; coxae with only short pale vestiture,
no strong setae; all tarsi with strong black claws and large
yellowish pulvilli; I: 6.7; 7.0; 3.6/ 1.2/ 1.0/ 0.9/ 0.8; FI and
TI mostly with short vestiture; TI without anteroapical
comb; TI, It1, and It2 each with 2 pairs of av-pv setae, and
It3 each with single pair of av-pv setae; II: 6.7; 7.0; 3.0/ 1.4/
1.0/ 0.9/ 0.7; TII with pair strong apical av-pv setae setae;
each tarsomere IIt1–3 with apical av-pv setae; IIt1 with black
Bickel: a new empidid fly genus
ventral pile; remainder of tarsus II with dense pale ventral
pile; III: 11.0; 10.0; 3.0/ 1.3/ 1.0/ 0.8/ 0.8; FIII gradually
expanding apically, and weakly clavate in distal third; TIII
without strong setae; IIIt with short black vestiture.
Abdomen entirely shining black, with metallic blue-violet
reflections; without pruinosity; tergum 7 with distal Ushaped excavation; tergum and sternum 8 unmodified;
hypopygium (Fig. 3e); male cercus divided into short basal
cercal plate and rather broad clasping cercus.
321
in October and November. This species shows strong sexual
dimorphism in mesonotal pattern: males have an extensive
black area (Fig. 2d), whereas females have an entirely
yellow mesonotum (Fig. 2e).
Female similar to male except as noted: face and clypeus
both wider; thorax including scutellum and postnotum
entirely yellow, without any black markings (Fig. 2e);
vestiture of tarsus I and II not as strongly developed;
oviscapt Fig. 3f.
Remarks. Bandella noorinbee is known from coastal and
lowland sites from the northern coast of New South Wales
to East Gippsland, Victoria. All specimens were collected
Figure 4. Bandella tasmanica: a, male postabdomen, left lateral. Bandella duvalli: b, male postabdomen, left
lateral; c, male postabdomen, dorsal. Bandella maxi: d, male postabdomen. Bandella costalis: e,. male right tibia
and tarsus II, dorsal. Legend: aed, aedeagus; cer (cl), clasping cercus; cer (pl), cercal plate; epan, epandrium; hyp,
hypandrium; tg, tergum; st, sternum.
322
Records of the Australian Museum (2002) Vol. 54
Bandella maxi n.sp.
Bandella duvalli n.sp.
Figs. 2g, 4d
Figs. 1, 2h, 2i, 4b, 4c
Type material. HOLOTYPE 웧 (K 173140), Queensland: Mt
Lewis, Mt Lewis Rd nr forestry hut, 1100 m, 10.xii.1974,
at light, M.S. Moulds (AMS).
Type material. HOLOTYPE 웧, Tasmania: 14 km SW by S
of Wilmot, 41°30'S 146°05'E, 31.i.1983, I.D. Naumann &
J.C. Cardale (ANIC).
Description (male): length: 10.0; wing: 7.5×2.2. Head:
post-cranium, vertex and frons shining black with no
pruinosity; ventral postcranium with scattered pale setae;
postorbitals reduced to short hairs, white ventrally, black
dorsally; face as wide as frons and covered with silvery
pruinosity; palp yellow with white setae, curved and slightly
clavate; proboscis yellow with dark brown apex, elongate,
length about one and a half head height, and projecting
ventrally; antenna dark brown.
Thorax: pleura and anterior mesonotum mostly redbrown; anterior mesonotum red, interdigitating with black
posterior mesonotum (Fig. 2g); scutellum black with reddish
margin; postnotum black, posterior slope of mesonotum
covered with field of short setulae.
Legs: coxae mostly reddish yellow; but CIII laterally
infuscated; coxae with only short pale vestiture, no strong
setae; all tarsi with strong black claws and large yellowish
pulvilli; I: 9.0; 9.8; 5.0/ 2.1/ 1.5/ 1.8/ 1.4; leg I mostly redyellow; FI and TI mostly with short vestiture; TI without
anteroapical comb; TI and It1 each with 2–3 pairs of apical
av and pv setae, It2 and It3 each with single av-pv setal pair;
all of tarsus I with pale ventral pile; II: 10.0; 10.5; 4.5/ 1.8/
1.3/ 1.4/ 1.0; FII mostly reddish yellow but infuscated in
distal half; TII and IIt1 red-yellow; distal tarsomeres II dark
brown; TII with pair strong av setae; each tarsomere IIt1–3
with apical av-pv setae; IIt3 and IIt4 with dense pale ventral
pile; III: 13.5; 13.0; 5.0/ 2.0/ 1.3/ 1.5/ 1.3; FIII yellowish in
basal half but distally dark brown; TIII yellow in basal third,
and with distal two-thirds of TIII, and all IIIt dark brown;
FIII gradually expanding apically, and weakly clavate in
distal third; TIII without strong setae; IIIt with short black
vestiture.
Abdomen: terga 1 and 2 reddish, remainder of abdomen
black; cuticle glabrous, without pruinosity; segment 7 with
well-developed tergum and sternum, tergum 7 not
excavated; nor differentiated from segments anteriad;
tergum 8 reduced to narrow band; sternum 8 unmodified;
hypopygium (Fig. 4d).
Additional material. Tasmania: 웧, Arthur Plains,
18.ii.1965; 웨, Mt Bowes, NE slope, 11.ii.1965 (MVM);
웧, Mt Wellington, Hobart, 42°55'S 147°14'E, 1.ii.1992, 720
m, on Leptospermum scoparia (UQIC); 웨, Cradle Valley,
26.i.1923; 웧, Strahan, 6.ii.1923; 웨, Mt Farrel, 9.ii.1923;
웧, Mt. Field NP, 950 m, 22.ii.1963 (ANIC); 웧, Mt Barrow,
41°22'S 147°25'E, 900 m, 12.ii1988 (AMS).
Female unknown.
Remarks. Bandella maxi is known only from submontane
rainforest above 1000 m on Mt. Lewis, in the Queensland
wet tropics. It has a greatly expanded and keel-like
hypopygium.
However, male abdominal segments 7 and 8 are relatively
unmodified, and therefore plesiomorphic with respect to
the other Bandella species. This species is named for the
collector, Max Moulds.
Description (male): length: 10.0; wing: 7.5×2.2 (habitus,
Fig. 1). Head: post-cranium black, covered laterally and
ventrally with grey pruinosity; dorsal post-cranium, vertex
and frons black without pruinosity; postcranium with usual
pale ventral hairs and with black dorsal hairs; postorbital
setae white ventrally, black dorsally; lateral frons with
setulae more distinct than usual; palp yellow with white
setae, curved and slightly clavate; labrum dark brown
basally, yellow distally; labellum dark brown; antenna black;
scape and pedicel with short black setulae.
Thorax: pleura mostly black but red-brown near sutures,
and covered with grey pruinosity; anterior mesonotum redbrown, interdigitating with black posterior mesonotum,
black lateral patches and black ac band, but in some
specimens this pattern is obscured (Fig. 2h,i); scutellum
red-brown with black base; postnotum black with grey
pruinosity; posterior slope of mesonotum covered with field
of short setulae.
Legs: coxae dark brown to black although CI and CII
reddish brown distally; coxae with only short pale vestiture,
no strong setae; all tarsi with strong black claws and large
yellowish pulvilli; legs I and II mostly red-yellow; It1–3,
each with pairs of av setae; each tarsomere IIt1–3 with some
subapical av setae; FIII yellowish in basal two thirds, but
dark brown and slightly clavate in distal third; TIII yellow
at base, dark brown from 1 3–5 6, and yellow in distal sixth;
distal sixth of TIII and IIIt 1 with pale yellow vestiture;
IIIt2–5 with black vestiture.
Abdomen: tergum 1 black; preabdomen translucent redbrown, with some specimens showing black areas laterally
and dorsally on terga 4 and 5; cuticle glabrous, without
pruinosity; tergum 7 only slightly excavated; tergum 8 with
lateral flaplike projections; sternum 8 unmodified;
hypopygium mostly black (Fig. 4b,c).
¤
¤
Female similar to male except as noted: frons wider, thorax
with black markings smaller in extent; leg and vestiture
colour similar; abdomen mostly orange in colour.
Remarks. Bandella duvalli is known from various locales
in western Tasmania, mostly in upland areas. There is some
intraspecific variation in the intensity of mesonotal
infuscation among the specimens, as shown in Fig. 4b and 4c.
This species named in honor of Steven G. Duvall of
California, who helped sponsor this research.
Bickel: a new empidid fly genus
323
Bandella tasmanica n.sp.
Bandella costalis n.sp.
Figs. 2j, 4a
Figs. 2k, 4e
Type material. HOLOTYPE, 웧, Tasmania: Pelion Gap, 2
km ENE of Mt Ossa, 41°52'S 146°03'E, 1120 m, 7.iii.1991–
9.iv.1991, malaise trap, E. Edwards & J. Berry, ANIC.
PARATYPES: 2웧, same data as holotype; 1웧, Pelion Hut, 3
km S of Mt Oakleigh, 41°50'S 146°03'E, 860 m, iii.1991,
malaise trap; 1웧, Pelion Hut, same data but 6–11.iii.1991,
pans (all ANIC).
Type material. HOLOTYPE, 웧, PARATYPE, 웨, Western
Australia: 11 km S of Pemberton, 34°32'S 116°02'E,
6.x.1981, I. Naumann & J. Cardale (both ANIC).
Additional material. Tasmania: 웨, Derwent Bridge,
12.ii.1967 (ANIC).
Description (male): length: 11.0–11.5; wing: 8.0×2.2;
similar to B. duvalli, except as noted. Head also with black
hairs on dorsal postcranium, scape and pedicel.
Thorax: pleura and postnotum black and covered with
dense grey pruinosity; anterior mesonotum mostly orange;
but with black ac band which continues and is slightly wider
across posterior slope to scutellum; large black spot laterad
of dc row and posteriad of mesonotal suture, and black pa
spot, which is sometimes only weakly developed (Fig. 2j);
postpronotum and postalar callus pale yellow; scutellum
orange but dark brown at base; dc band with short setulae
extending to scutellum, and with some short setulae laterally.
Legs: coxae mostly reddish yellow; but CII and CIII
infuscated at very base; trochanters yellow with dark brown
apices; all femora and tibiae reddish yellow, except TIII
with brown band from 3 5 to 3 4 which varies somewhat in
extent; It brownish, IIt mostly yellow, IIIt yellow; coxae
with only short pale vestiture, no strong setae; all tarsi curled
in repose; It1–3 each with pairs of ad-pd setae; IIIt1 ventrally
with dense pale pile and some short dark setulae.
Abdomen: tergum 1 black; segments 2–6 entirely orangeyellow; tergum 8 also with lateral flaplike projections;
hypopygium (Fig. 4a) mostly yellow except for dark brown
apex and edge of hypandrial keel and apices of epandrium
and cercus.
¤
¤
Female. Putative female specimen similar to male except
as noted: thoracic pleura with mixed black and yellow
patches, and with distal postnotum black; mesonotum
entirely orange with only dark brown ac stripe present,
thinner than that of male; leg colour similar and TIII also
with brown band.
Remarks. Bandella tasmanica is known only from the
Central Highlands of Tasmania, and most specimens were
collected near Mt. Pelion. This species is close to B. duvalli,
but the two species can be readily separated by the set of
diagnostic key characters: the pattern of the mesonotum,
and colour of coxae and abdomen. Both species have a
similar banded tibia III and have a short flap like tergum 8,
that rests against the hypopygium.
Additional material: Western Australia: 3웧, 3웨, 10 km N
of Walpole, 7.x.1970; 7웧, 6웨, Nornalup NP, 9.x.1970, coll.
D.H. Colless, ANIC.
Description (male): length: 10.0; wing: 7.5×2.2. Head:
vertex and frons shining black with no pruinosity; ventral
postcranium with scattered pale setae; postorbitals as short
hairs, white ventrally, black dorsally; face covered with
silvery pruinosity; labrum yellow, labellum dark brown;
antenna dark brown; scape and pedicel setose.
Thorax: pleura black, although meron and laterotergite
with large yellowish areas, and covered with grey pruinosity;
mesonotum and scutellum entirely black and shining with
no pruinosity (Fig. 2k); postnotum black but covered with
grey pruinosity; entire mesonotum with only setulae, no
major setae present; ac band narrow, 2–3 setulae wide, with
bare cuticle each side of this band extending to posterior
slope, and with narrow dc row.
Legs: coxae mostly reddish yellow; remainder of legs
yellow, except apex of trochanters and very base of femora
black, and distalmost tarsomeres infuscated; coxae with only
short pale vestiture, no strong setae, remainder of legs with
black vestiture; TI with pair apical av-pv setae, It1 with very
long posterior setae, and with slightly longer than normal
anterior setae; It2–3 each with 2–3 pairs of long anterior and
posterior setae (MSSC); TII (Fig. 4e) in distal half with
some long anterior and posterior setae, and apically with
some strong av, pv and ventral setae (MSSC); IIt1 with very
long anterior and posterior setae, and It2–3 with long posterior
setae only (both MSSC); IIt4–5 unmodified (MSSC); FIII
slightly clavate in distal half; TIII with some irregular
long setae, especially ventrally (MSSC); IIIt1 with long
posterior setae (MSSC), but not as long as those of IIt1;
IIIt2 with 3–4 long posterior setae, and IIIt3 with some long
apical setae (MSSC); IIIt4–5 unmodified.
Wing: stigma brownish and diffuse.
Abdomen: terga yellowish with dark-brown to black
posterior borders, and with weak dorsal infuscation;
postabdomen (not figured) with tergum 8 developed into
lateral flap-like projection; hypopygium mostly yellow, with
somewhat enlarged hypandrial keel which projects
posteriorly.
Female similar to male except as noted: face slightly wider;
tibiae and tarsi with only short unmodified vestiture, without
long anterior and/or posterior setae.
Remarks. Bandella costalis is known from wet sclerophyll
karri forests in southwestern Western Australia. Male tibiae
and tarsi variously have long anterior and/or posterior setae,
especially well developed on leg II (Fig. 4e).
324
Records of the Australian Museum (2002) Vol. 54
Bandella albitarsis n.sp.
Figs. 2f
Type material. HOLOTYPE, 웧, New South Wales: New
England NP, 13.x.1962, D.H. Colless (ANIC).
Additional material. New South Wales: 웨, Dorrigo NP,
15.x.1966 (ANIC); 웧, Blue Mountains, Blackheath, x.1930
(MVM).
Description (male): length: 8.0; wing: 7.5×2.2. Head: postcranium shining black, with silvery pruinose band around
outer rim except near vertex; frons shining black with no
pruinosity; ventral postcranium with scattered pale setae;
postorbitals entirely white, within silvery pruinose band;
face covered with golden pruinosity; labrum and labellum
yellow; scape and pedicel yellow-brown; first flagellomere
dark brown.
Thorax: cuticle glabrous with little pruinosity; pleura and
mesonotum mostly red-brown, but mesonotum with black
areas laterally, and with pale yellow humeral, pronotal and
notopleural areas (Fig. 2f); scutellum orange; postnotum
mostly dark brown; distinct patches of orientated silvery
pruinosity visible in posterolateral view on following
thoracic areas: middle of postnotum, anatergite, meron, and
area of anepisternum just anteriad of wind base; posterior
slope of mesonotum with scattered short setulae.
Legs: coxae, trochanters and all femora reddish yellow
except trochanters with black apices; leg vestiture mostly
yellowish; coxae with only short vestiture, no strong setae;
all tarsi with strong black claws and large yellowish pulvilli;
TI and It1 dark brown to black with reddish vestiture, in
sharp contrast to the ivory white It2–5; TI and each tarsomere
It1–3 with some short av-pv setae; TI without distinct
anteroapical comb; It1 slightly swollen (MSSC); TII and
tarsus II red yellow; TII with ad row of 8–10 short yellowish
setae along length; IIt1–3 each with some short av-pv setae;
IIt3 and IIt4 with dense pale ventral pile; TIII and IIIt1 red
yellow, and IIIt2–5 distinctly pale yellow; FIII not clavate;
IIIt with short vestiture.
Abdomen entirely red-brown, with some brownish
infuscation distally; cuticle glabrous, without pruinosity;
tergum 7 slightly excavated distally; hypopygium (not
figured) almost entirely yellow.
Female similar to male except as noted: post-cranium also
with silvery pruinose band around outer rim; thorax of
similar colour and with similar pruinose patches; TI
yellowish basally but dark brown in distal half; It1 black
but not swollen; tarsomeres t2–5,on all three legs distinctly
white; leg vestiture somewhat weaker than on male.
Remarks. Bandella albitarsis is known from the Dorrigo
Escarpment in northeastern New South Wales, and the Blue
Mountains west of Sydney, both at elevations above 750
m. All specimens were collected in October.
This species has several diagnostic characters, developed
in both sexes: (a) Bright silvery pruinose areas are distinctly
visible in oblique view on the following areas: a band around
the rim of the postcranium, middle of the postnotum, the
anatergite, most of the meron, and the area of anepisternum
just anteriad of wing base. These silvery patches may
facilitate mating in flight, especially since incident light
would make them appear to “flash” while turning. (b) Leg
I has tarsomeres 2–5 ivory white in both sexes. In males,
tarsi II and III are mostly yellow, whereas in females, all
tarsomeres 2–5 on legs III and III are distinctly white. The
white distal tarsomeres are more strongly developed on
females than males, and should be regarded as a female
secondary sexual character (FSSC) that is only weakly
developed in males.
The male from New England National Park had
Leptospermum pollen on its proboscis, and probably was
nectar feeding.
ACKNOWLEDGMENTS. I thank the following institutions and their
curators for loan of specimens: AMS, Australian Museum, Sydney,
M. Moulds; ANIC, Australian National Insect Collection, CSIRO,
Canberra, P. Cranston; CNC, Canadian National Collection,
Ottawa, J. Cumming; MVM, Museum of Victoria, Melbourne, K.
Walker. B. Sinclair provided both sound advice and specimens
(deposited CNC) for this project. S. Bullock drew the genitalic
figures. The habitus of Bandella duvalli was drawn by H. Finlay.
This research was supported by Australian Research Council grant
A19232651, for study of the systematics of Australian Empidinae.
Additional support was provided by S. Duvall through the
Australian Museum “Immortals” programme.
References
Bickel, D.J., 1996. Thinempis, a new genus from Australia and
New Zealand (Diptera: Empididae), with notes on the tribal
classification of the Empidinae. Systematic Entomology 21:
115–128.
Bickel, D.J., 1998. Cunomyia, a distinctive new hilarine fly
genus from the Tasmanian World Heritage Area (Diptera:
Empididae). Papers and Proceedings of the Royal Society
of Tasmania 132: 59–63.
Chvála, M., 1976. Swarming, mating, and feeding habits in
Empidinae (Diptera), and their significance in evolution of the
family. Acta Entomologica Bohemoslovaca 73: 353–366.
Chvála, M., 1983. The Empidoidea (Diptera) of Fennoscandia
and Denmark II. General Part. The Families Hybotidae,
Atelestidae, and Microphoridae. Fauna Entomologica
Scandinavica 12: 1–279.
Collin, J.E., 1928. New Zealand Empididae. London: British
Museum (Natural History), pp. 110.
Collin, J.E., 1933. Empididae, Part 4, in Diptera of Patagonia
and South Chile. London: British Museum (Natural History),
pp. 334.
Cumming, J.M., B.J. Sinclair & D.M. Wood, 1995. Homology
and phylogenetic implications of male genitalia in Diptera—
Eremoneura. Entomologica scandinavica 26: 120–151.
Gauld, I., & L.A. Mound, 1982. Homoplasy and the delineation
of holophyletic genera in some insect groups. Systematic
Entomology 7: 73–86.
McAlpine, J.F., 1981. Morphology and terminology. In Manual
of Nearctic Diptera, ed. J.F. McAlpine et al., vol 1, pp. 9–64.
Ottawa: Agriculture Canada.
Sinclair, B.J., 1995. Generic revision of the Clinocerinae
(Empididae), and description and phylogenetic relationships
of the Trichopezinae, new status (Diptera: Empidoidea).
Canadian Entomologist 127: 665–752.
Smith, K.G.V., 1969. The Empididae of southern Africa (Diptera).
Annals of the Natal Museum 19: 1–342.
Manuscript received 23 April 2001, revised 11 October 2001 and accepted
29 October 2001.
Associate Editor: S.F. McEvey